Article type: Letter 41 42 One Sentence Summary: Marine heatwaves alter ecosystem structure and functioning at 43 global scales. 44 45 46 47 48 49 implications for marine ecosystems 1 . Concurrent with long-term persistent warming, 50 discrete periods of extreme regional ocean warming (marine heatwaves, 'MHWs') have 51 increased in frequency 2 . Here we quantify trends and attributes of MHWs across all 52 ocean basins and examine their biological impacts from species to ecosystems. Multiple 53 regions within the Pacific, Atlantic and Indian Oceans are particularly vulnerable to 54 MHW intensification, due to the co-existence of high levels of biodiversity, a prevalence 55 of species found at their warm range edges, or concurrent non-climatic human impacts. 56 The physical attributes of prominent MHWs varied considerably, but all had 57 deleterious impacts across a range of biological processes and taxa, including critical 58 foundation species (corals, seagrasses and kelps). MHWs, which will likely intensify 59 with anthropogenic climate change 3 , are rapidly emerging as forceful agents of 60 disturbance with the capacity to restructure entire ecosystems and disrupt the provision 61 of ecological goods and services in coming decades. 62 63 Anthropogenic climate change is driving the redistribution of species and reorganization of 64 natural systems and represents a major threat to global biodiversity 4,5 . The biosphere has 65 401 working group on marine heatwaves (www.marineheatwaves.org).

Ocean acidification and warming are considered two of the greatest threats to marine biodiversity, yet the combined effect of these stressors on marine organisms remains largely unclear. Using a meta-analytical approach, we assessed the biological responses of marine organisms to the effects of ocean acidification and warming in isolation and combination. As expected biological responses varied across taxonomic groups, life-history stages, and trophic levels, but importantly, combining stressors generally exhibited a stronger biological (either positive or negative) effect. Using a subset of orthogonal studies, we show that four of five of the biological responses measured (calcification, photosynthesis, reproduction, and survival, but not growth) interacted synergistically when warming and acidification were combined. The observed synergisms between interacting stressors suggest that care must be made in making inferences from single-stressor studies. Our findings clearly have implications for the development of adaptive management strategies particularly given that the frequency of stressors interacting in marine systems will be likely to intensify in the future. There is now an urgent need to move toward more robust, holistic, and ecologically realistic climate change experiments that incorporate interactions. Without them accurate predictions about the likely deleterious impacts to marine biodiversity and ecosystem functioning over the next century will not be possible.